In many wild species, seeds are dormancy at maturity and will not germinate even under favorable environment conditions. Dormancy is a complex trail that is determined by many factors. Some studies have shown that cold stratification and the application of gibberellic acid (GA) can break seed dormancy and promote seed germination. The present study investigated the causes of plant dormancy and the effect of cold stratification and different concentrations of exogenous GA 3 in regulating Cephalotaxus sinensis seed germination. Results showed that C. Sinensis seeds have good water permeability, which suggested that seed coats were not the main cause that inhibited the seed germination. There were germination inhibitions in all parts of seeds, and the order of inhibitory effect was: testa < endosperm (embryo), which indicated that existence of germination inhibitions was the main reason causing seed dormancy. Endogenous GAs and IAA (indole-3-acetic acid) content increased, while ABA (abscisic acid) content decreased over the experiments. ZR (zeatin riboside) content decreased in the early phase of cold stratification, but rebounded by the end of the experimental period. The changes of endogenous hormone indicated that GA, IAA and ZR played a positive role in seed germination, whereas ABA was associated with seed dormancy. Besides, the relative ratio of GA/ABA, IAA/ABA and ZR/ABA may play a more important role than their absolute level during the seed development.
Nasreen, S., Yousaf, M., Mohmand, A.S. and Mailk, M.A. (2012) Study of Seed Dormancy Mechanisms; Causes and Control. Asian Journal of Plant Sciences, 1, 210-212.
Olmez, Z., Temel, F., Gokturk, A. and Yahyaoglu, Z. (2007) Effects of Sulphuric Acid and Cold Stratification Pretreatments on Germination of Pomegranate (Punica granatum L.) Seeds. Asian Journal of Plant Sciences, 6, 427-430. https://doi.org/10.3923/ajps.2007.427.430
Yang, Q.H., Ye, W.H., Song, S.Q. and Yin, S.H. (2003) Summarization on Cause of Seed Dormancy and Dormancy Polymorphism. Acta Botanica Boreali-Occidentalia Sinica, 23, 837-843.
Pawlowski, T.A. (2009) Proteome Analysis of Norway Maple (Acer platanoides L.) Seeds Dormancy Breaking and Germination: Influence of Abscisic and Gibberellic Acids. BMC Plant Biology, 9, 1-13.
Black, M., Bewley, J.D. and Halmer, P. (2008) The Encyclopedia of Seeds: Science, Technology and Uses. CAB International, Wallingford.
Koornneef, M., Bentsink, L. and Hilhorst, H. (2002) Seed Dormancy and Germination. Current Opinion in Plant Biology, 5, 33-36. https://doi.org/10.1016/S1369-5266(01)00219-9
Carlón, L. (1998) Distribución geográfica y comportamiento ecológico de Helianthemum tinetense Mayor López & Fernández Benito, endemismoastur-galaico. Bol. Ciencias Nat. RIDEA, 45, 1-12.
López González, G. (1993) Helianthemum Mill. In: Castroviejo, S., Aedo, C., Cirujano, S., Laínz, M., Montserrat, P., Morales, R., Munoz Garmendia, F., Navarro, C., Paiva, J. and Soriano, C., Eds., Flora Iberica 3, Real Jardín Botánico—CSIC, Madrid, 365-421.
Perez-Garca, F. (2006) Seed Germination of Five Helianthemum Species: Effect of Temperature and Presowing Treatments. Journal of Arid Environments, 65, 688-693. https://doi.org/10.1016/j.jaridenv.2005.10.008
Khan, A.A. (1977) The Physiology and Biochemistry of Seed Dormancy and Germination. North Horth Holland Publishing Company, Amsterdam, 1-75.
Hilhorst, H.W.M. (1995) A Critical Update on Seed Dormancy. I. Primary Dormancy. Seed Science Research, 5, 61-73. https://doi.org/10.1017/S0960258500002634
Hidayati, S.N., Baskin, J.M. and Baskin, C.C. (2000) Dormancy-Breaking and Germination Requirements of Seeds of Four Lonicera Species (Caprifoliaceae) with Underdeveloped Spatulate Embryos. Seed Science Research, 10, 459-469.
Ma, S. (2006) Study on the Dormancy Mechanism of Acanthopanax senticosus Seeds. Jilin Agricultural University, Jilin.
Finch-Savagel, W.E. and Leubner-Metzger, G. (2006) Seed Dormancy and the Control of Germination. New Phytologist, 171, 501-523. https://doi.org/10.1111/j.1469-8137.2006.01787.x
Baskin, J.M. and Baskin, C.C. (2008) Some Considerations for Adoption of Nikolaeva’s Formula System into Seed Dormancy Classification. Seed Science Research, 18, 131-137. https://doi.org/10.1017/S096025850803674X
Benech-Arnold, R.L.M. and Giallorenzic, F.J. (1999) Termination of Hull-Imposed Dormancy in Developing Barley Grains Is Correlated with Changes in Embryonic ABA Levels and Sensitivity. Seed Science Research, 9, 39-47. https://doi.org/10.1017/S0960258599000045
Zhang, L.P., Chen, Z. and Li, X.N. (1999) Seed Germination Characteristics and Embryo Development of Schisandra chinensis (Turcz.) Bail. Journal of Chinese Material Medica, 24, 459-461.
Zhang, X.W., Cao, J.J., Gong, Y.X. and Zhang, X.P. (2007) Seed Dormancy and Germination in Rare Plant Pteroceltis tatarinowii Maxim. Journal of Biology, 24, 28-31.
Arditti, J. and Pray, T.R. (1969) Dormancy Factors in Iris (Iridaceae) Seeds. American Journal of Botany, 56, 254-259. https://doi.org/10.2307/2440847
Yagihashi, T., Hayashida, M. and Miyamoto, T. (1998) Effects of Bird Ingestion on Seed Germination of Sorbus commixta. Oeco1ogia, 114, 209-212.
Paulsen, T.R. and Hogstedt, G. (2002) Passage through Bird Guts Increases Germination Rate and Seedling Growth in Sorbus aucuparia. Functional Ecology, 16, 608-616. https://doi.org/10.1046/j.1365-2435.2002.00668.x
Randolph, L.F. and Cox, L.G. (1943) Factors Influencing the Germination of Iris Seed and the Relation of Inhibiting Substances to Embryo Dormancy. Proceedings of the American Society for Horticultural Science, 43, 284-300.
Khan, A.A. (1975) Primary, Preventive and Permissive Role of Hormones in Plant Systems. The Botanical Review, 41, 391-420. https://doi.org/10.1007/BF02860831
Kucera, B., Cohn, M.A. and Leubner-Metzger, G. (2005) Plant Hormone Interaction during Seed Dormancy Release and Germination. Seed Science Research, 15, 281-307. https://doi.org/10.1079/SSR2005218
Hu, X.W., Huang, X.H. and Wang, Y.R. (2012) Hormonal and Temperature Regulation of Seed Dormancy and Germination in Leymus chinensis. Plant Growth Regulation, 67, 199-207. https://doi.org/10.1007/s10725-012-9677-3
Abdallah, A., Debez, A., Barhoumi, Z., Smaoui, A. and Abdelly, C. (2009) ABA, GA3, and Nitrate May Control Seed Germination of Crithmum maritimum (Apiaceae) under Saline Conditions. Comptes Rendus Biologies, 332, 704-710. https://doi.org/10.1016/j.crvi.2009.03.009
Steffens, B., Wang, J. and Sauter, M. (2006) Interactions between Ethylene, Gibberellin and Abscisic Acid Regulate Emergence and Growth Rate of Adventitious Roots in Deepwater Rice. Planta, 223, 604-612. https://doi.org/10.1007/s00425-005-0111-1
Le Page-Degivry, M.T., Bianco, J., Barthe, P. and Garello, G. (1996) Change in Hormone Sensitivity in Relation to Onset and Breaking of Sunflower Embryo Dormancy. In: Lang, G.A., Ed., Plant Dormancy: Physiology, Biochemistry and Molecular Biology, CAB International, Wallingford, 221-231.
Grappin, P., Bouinot, D., Sotta, B., Miginiac, E. and Jullien, M. (2000) Control of Seed Dormancy in Nicotiana plumbaginifolia: Post-Imbibition Abscisic Acid Synthesis Imposes Dormancy Maintenance. Planta, 210, 279-285. https://doi.org/10.1007/PL00008135
Christmann, A., Moes, D., Himmelbach, A., Yang, Y., Tang, Y. and Grill, E. (2006) Integration of Abscisic Acid Signaling into Plant Responses. Plant Biology, 8, 314-325. https://doi.org/10.1055/s-2006-924120
Baskin, J.M. and Baskin, C.C. (2004) A Classification System for Seed Dormancy. Seed Science Research, 14, 1-16. https://doi.org/10.1079/SSR2003150
Chen, S.Y., Chen, C.T., Baskin, J.M. and Baskin, C.C. (2010) Storage Behavior and Changes in Concentrations of Abscisic Acid and Gibberellins during Dormancy Break and Germination in Seeds of Phellodendron amurense var. wilsonii (Rutaceae). Tree Physiology, 30, 275-281. https://doi.org/10.1093/treephys/tpp111
Lewak, S. (2011) Metabolic Control of Embryonic Dormancy in Apple Seed: Seven Decades of Research. Acta Physiologiae Plantarum, 33, 1-24. https://doi.org/10.1007/s11738-010-0524-8
Benech-Arnold, R.L., Steinbach, H.S., Kristof, G. and Sa’nchez, R.A. (1995) Fluctuating Temperatures Have Different Effects on Embryonic Sensitivity to ABA in Sorghum Varieties with Contrasting Pre-Harvest Susceptibility. Journal of Experimental Botany, 46, 711-717. https://doi.org/10.1093/jxb/46.6.711
Chien, C.T., Kuo-Huang, L.L. and Lin, T.P. (1998) Changes in Ultrastructure and Abscisic Acid Level, and Response to Applied Gibberellins in Taxus mairei Seeds Treated with Warm and Cold Stratification. Annals of Botany, 81, 41-47. https://doi.org/10.1006/anbo.1997.0542
Singh, Z. and Browning, G. (1991) The Role of ABA in the Control of Apple Seed Dormancy Reappraised by Combined Gas Chromatography Mass Spectrometry. Journal of Experimental Botany, 42, 269-275. https://doi.org/10.1093/jxb/42.2.269
Kochanko, V.G., Gitzesik, M., Chojnowsk, M., et al. (1998) Effect of Temperature, Growth Regulators and Other Chemicals on Echinacea purpurea (L.) Moench Seed Germination and Seedling Survival. Seed Science and Technology, 26, 547-554.
Yang, W.X. and Fang, S.Z. (2008) Dynamic Changes of Endogenous Hormones in Cyclocarya paliurus Seed during Stratification. Journal of Nanjing Forestry University (Natural Sciences Edition), 35, 85-88.
Li, K.J., Jin, Y.J., Chen, H.J., et al. (1997) Changes of Endogenous GAs and IAA Contents in Pinus koraiensis Seeds during Cold Stratification and Germination. Hebei Journal of Forestry and Orchard Research, 12, 203-208.
Wang, S.G. (1999) Cytokinins and Plant Seed Development and Germination. Seed, 105, 35-37.
Cao, B.H. and Cai, C.J. (2006) Study on the After-Ripening Physiology and Endogenous Hormones of Ginkgo biloba Seeds. Scientia Silvae Sinicae, 42, 32-37.
Liu, X.J., Wang, D., Yu, D.L. and Liang, P. (2013) Introduction Performance and Breeding Technology of Cephalotaxus sinensis in Xiongyue Area. Northern Horticulture, 8, 85-87.
Zhou, X.J., Hu, Z.B., Huang, L.D. and Zhang, G.Y. (1997) Studies on Cephalotaxus Plants Resources over China. Journal of Hubei Agricultural College, 17, 100-103.
Bocar, M., Jossang, A. and Bodo, B. (2003) New Alkaloids from Cephalotaxus fortunei. Journal of Natural Products, 66, 152-154. https://doi.org/10.1021/np0203178
Muhammad, K.S., Deng, Y.L. and Dai, R.J. (2008) Attenuation of Biochemical Parameters in Streptozotocin-Induced Diabetic Rats by Oral Administration of Extracts and Fractions of Cephalotaxus sinensis. Journal of Clinical Biochemistry and Nutrition, 42, 21-28.
Hao, S.H., Wei, Y., Zhang, J. and Zhang, X. (2006) Herbicidal Activity of Twig and Needle Extracts from Cephalotaxus sinensis against Amaranthus retroflexus. Chinese Journal of Pesticide Science, 8, 91-94.
Shi, H., An, S.P., Yang, W., Feng, X.Q., Wang, M.Q. and Zhang, J.T. (2007) Study on Cutting and Breeding Trials of Cephalotaxus chinensis. Journal of Gansu Forestry Science and Technology, 32, 23-25.
Pang, X.H. and Song, J.Y. (2010) Research Progress of Resources in Cephalotaxus Mannii. Shanxi Journal of Agricultural Sciences, 1, 119-121.
Wang, X.P. and Wang, J.L. (1998) Extraction, Separation and Biological Measurement of Endogenous Growth Regulating Substance of Seeds of Pinus bungeana. Seeds, 5, 19-23.
Zeng, Q.Q., Chen, H.B., Lu, C.H. and Li, J.G. (2006) An Optimized HPLC Procedure for Analyzing Endogenous Hormones in Different Organs of Litchi. Journal of Fruit Science, 23, 145-148.
Yan, S., et al. (2014) Optimization of Seed Germination and Detection of Germination Inhibitors in Gardenia Fruits. Journal of Southern Agriculture, 45, 376-382.
Linkies, A., Graeber, K., Knight, C. and Leubner-Metzger, G. (2010) The Evolution of Seeds. New Phytologist, 186, 817-831. https://doi.org/10.1111/j.1469-8137.2010.03249.x
Debeaujon, I., Léon-Kloosterziel, K.M. and Koornneef. M. (2000) Influence of the Testa on Seed Dormancy, Germination, and Longevity in Arabidopsis. Plant Physiology, 122, 403-413. https://doi.org/10.1104/pp.122.2.403
Greipsson, S. (2001) Effects of Stratification and GA3 on Seed Germination of a Sand Stabilizing Grass Leymus arenarius Used in Reclamation. Seed Science & Technology, 29, 1-10.
Yang, W.X., Fu, X.X. and Fang, S.Z. (2005) The Seed Coat Structure of Cyclocarya paliurys and Its Effects on Water Permeability. Journal of Nanjing Forestry University (Natural Sciences Edition), 29, 25-28.
Zhang, F.J., Xu, X.Y., Meng, X.D., Wang, F.B. and Jin, Y.J. (2004) Breaking the Seed Dormancy and Stimulating the Seed Germination of Gleditsia sinensis. Journal of Fujian College of Forestry, 24, 175-178.
Chen, J.L. (1994) A Study on the Seed Dormancy of Euonymus carnosus and Its Commutation. Forest Research, 7, 227-229.
Asakawa, S. (1956) Studies on the Delayed Germination of Fraxinus mandshurica var. Japonica Seeds. Bulletin of the Government Forest Experiment Station, 87, 75-84.
Graeber, K., Nakabayashi, K., Miatton, E., Leubner-Metzger, G. and Soppe, W.J.J. (2012) Molecular Mechanisms of Seed Dormancy. Plant, Cell and Environment, 35, 1769-1786. https://doi.org/10.1111/j.1365-3040.2012.02542.x
Evenari, M. (1949) Germination Inhibitors. The Botanical Review, 15, 153-194．
Chen, S.Y., et al. (2015) Effects of Moist Cold Stratification on Germination, Plant Growth Regulators, Metabolites and Embryo Ultrastructure in Seeds of Acer morrisonense (Sapindaceae). Plant Physiology and Biochemistry, 94, 165-173. https://doi.org/10.1016/j.plaphy.2015.06.004
Zhang, C.H., Zheng, Y.Q., Wu, J., Chen, P. and Li, B.J. (2012) Mechanisms of Seed Dormancy of Acer griseum. Bulletin of Botanical Research, 32, 573-577.
Sun, N., Chen, L., Liu, X.J. and Xing, L. (2016) The Effects of Germination Inhibitors Contained in Torreya grandis Seeds on Seed Germination. Journal of Huangshan University, 18, 44-47.
Huarte, H.R. and Benech-Arnold, R.L. (2010) Hormonal Nature of Seed Responses to Fluctuating Temperatures in Cynara cardunculus (L.). Seed Science Research, 20, 39-45. https://doi.org/10.1017/S0960258509990249
Leubner-Metzger, G. (2003) Functions and Regulation of β-1,3-Glucanase during Seed Germination, Dormancy Release and After-Ripening. Seed Science Research, 13, 17-34. https://doi.org/10.1079/SSR2002121
Nambara, E. and Marion, P.A. (2010) ABA Action and Interactions in Seeds. Trends in Plant Science, 8, 213-217. https://doi.org/10.1016/S1360-1385(03)00060-8
Ye, Y.H., Zhang, W.X., Zhang, S.W., Huang, H.L. and Ma, Y.X. (2014) The Effects of Cold Stratification on Dormancy Breaking and Endogenous Hormones Changes in Seeds of Sinopodophyllum hexandrum. Li Shizhen Medicine and Material Medica Research, 25, 715-717.
Wu, J., Li, Y.G., Luo, X.B., Tang, Y.Q., Zhou, Y. and Liu, X.H. (2013) Dynamic Changes of Endogenous Hormones in Styrax tonkinensis Seed during the Treatment Process. Acta Agriculturae Universitatis Jiangxiensis, 35, 988-992.
Gao, C.H. (2016) Effects of Exogenous GA on the Endogenous Hormones Content of Acanthopanax senticosus Harms in Ripening. Seeds, 35, 35-37.
Chen, W., Yang, Y., Ma, S.B. and Li, J. (2015) Study on Dormancy Type of Paris polyphylla var. yunnanensis Seeds. Southwest China Journal of Agricultural Sciences, 28, 783-786.
Zhao, X., Ma, X.J., Kaisar, Su, L.M. and Shi, J. (2006) Rule of Breaking Ferula fukanensis Seed Dormancy under Low-Temperature and Content Changes of Endogenous Hormone. Chinese Traditional and Herbal Drugs, 37, 268-270.